The Evolution of Empathy

(Editor’s Note: we are pleased to bring you this arti­cle thanks to our col­lab­o­ra­tion with Greater Good Mag­a­zine).

The Evo­lu­tion of Empathy

Empa­thy’s not a unique­ly human trait, explains pri­ma­tol­o­gist Frans de Waal. Apes and oth­er ani­mals feel it as well, sug­gest­ing that empa­thy is tru­ly an essen­tial part of who we are.

Once upon a time, the Unit­ed States had a pres­i­dent known for a pecu­liar facial dis­play. In an act of con­trolled emo­tion, he would bite his low­er lip and tell his audi­ence, “I feel your pain.” Whether the dis­play was sin­cere is not the issue here; how we are affect­ed by anoth­er’s predica­ment is. Empa­thy is sec­ond nature to us, so much so that any­one devoid of it strikes us as dan­ger­ous or men­tal­ly ill.

At the movies, we can’t help but get inside the skin of the char­ac­ters on the screen. We despair when their gigan­tic ship sinks; we exult when they final­ly stare into the eyes of a long-lost lover.

We are so used to empa­thy that we take it for grant­ed, yet it is essen­tial to human soci­ety as we know it. Our moral­i­ty depends on it: How could any­one be expect­ed to fol­low the gold­en rule with­out the capac­i­ty to men­tal­ly trade places with a fel­low human being? It is log­i­cal to assume that this capac­i­ty came first, giv­ing rise to the gold­en rule itself. The act of per­spec­tive-tak­ing is summed up by one of the most endur­ing def­i­n­i­tions of empa­thy that we have, for­mu­lat­ed by Adam Smith as “chang­ing places in fan­cy with the sufferer.”

Even Smith, the father of eco­nom­ics, best known for empha­siz­ing self-inter­est as the lifeblood of human econ­o­my, under­stood that the con­cepts of self-inter­est and empa­thy don’t con­flict. Empa­thy makes us reach out to oth­ers, first just emo­tion­al­ly, but lat­er in life also by under­stand­ing their situation.

This capac­i­ty like­ly evolved because it served our ances­tors’ sur­vival in two ways. First, like every mam­mal, we need to be sen­si­tive to the needs of our off­spring. Sec­ond, our species depends on coop­er­a­tion, which means that we do bet­ter if we are sur­round­ed by healthy, capa­ble group mates. Tak­ing care of them is just a mat­ter of enlight­ened self-interest.

Ani­mal empathy

It is hard to imag­ine that empathy—a char­ac­ter­is­tic so basic to the human species that it emerges ear­ly in life, and is accom­pa­nied by strong phys­i­o­log­i­cal reactions—came into exis­tence only when our lin­eage split off from that of the apes. It must be far old­er than that. Exam­ples of empa­thy in oth­er ani­mals would sug­gest a long evo­lu­tion­ary his­to­ry to this capac­i­ty in humans.

Evo­lu­tion rarely throws any­thing out. Instead, struc­tures are trans­formed, mod­i­fied, co-opt­ed for oth­er func­tions, or tweaked in anoth­er direc­tion. The frontal fins of fish became the front limbs of land ani­mals, which over time turned into hoofs, paws, wings, and hands. Occa­sion­al­ly, a struc­ture los­es all func­tion and becomes super­flu­ous, but this is a grad­ual process, and traits rarely dis­ap­pear alto­geth­er. Thus, we find tiny ves­tiges of leg bones under the skin of whales and rem­nants of a pelvis in snakes.

Over the last sev­er­al decades, we’ve seen increas­ing evi­dence of empa­thy in oth­er species. One piece of evi­dence came unin­ten­tion­al­ly out of a study on human devel­op­ment. Car­olyn Zahn-Waxler, a research psy­chol­o­gist at the Nation­al Insti­tute of Men­tal Health, vis­it­ed peo­ple’s homes to find out how young chil­dren respond to fam­i­ly mem­bers’ emo­tions. She instruct­ed peo­ple to pre­tend to sob, cry, or choke, and found that some house­hold pets seemed as wor­ried as the chil­dren were by the feigned dis­tress of the fam­i­ly mem­bers. The pets hov­ered near­by and put their heads in their own­ers’ laps.

But per­haps the most com­pelling evi­dence for the strength of ani­mal empa­thy came from a group of psy­chi­a­trists led by Jules Masser­man at North­west­ern Uni­ver­si­ty. The researchers report­ed in 1964 in the Amer­i­can Jour­nal of Psy­chi­a­try that rhe­sus mon­keys refused to pull a chain that deliv­ered food to them­selves if doing so gave a shock to a com­pan­ion. One mon­key stopped pulling the chain for 12 days after wit­ness­ing anoth­er mon­key receive a shock. Those pri­mates were lit­er­al­ly starv­ing them­selves to avoid shock­ing anoth­er animal.

The anthro­poid apes, our clos­est rel­a­tives, are even more remark­able. In 1925, Robert Yerkes report­ed how his bonobo, Prince Chim, was so extra­or­di­nar­i­ly con­cerned and pro­tec­tive toward his sick­ly chim­panzee com­pan­ion, Panzee, that the sci­en­tif­ic estab­lish­ment might not accept his claims: “If I were to tell of his altru­is­tic and obvi­ous­ly sym­pa­thet­ic behav­ior towards Panzee, I should be sus­pect­ed of ide­al­iz­ing an ape.”

Nadia Lady­gi­na-Kohts, a pri­ma­to­log­i­cal pio­neer, noticed sim­i­lar empath­ic ten­den­cies in her young chim­panzee, Joni, whom she raised at the begin­ning of the last cen­tu­ry, in Moscow. Kohts, who ana­lyzed Joni’s behav­ior in the minut­est detail, dis­cov­ered that the only way to get him off the roof of her house after an escape—much more effec­tive than any reward or threat of punishment—was by arous­ing sympathy:

If I pre­tend to be cry­ing, close my eyes and weep, Joni imme­di­ate­ly stops his plays or any oth­er activ­i­ties, quick­ly runs over to me, all excit­ed and shagged, from the most remote places in the house, such as the roof or the ceil­ing of his cage, from where I could not dri­ve him down despite my per­sis­tent calls and entreaties. He hasti­ly runs around me, as if look­ing for the offend­er; look­ing at my face, he ten­der­ly takes my chin in his palm, light­ly touch­es my face with his fin­ger, as though try­ing to under­stand what is hap­pen­ing, and turns around, clench­ing his toes into firm fists.

These obser­va­tions sug­gest that apart from emo­tion­al con­nect­ed­ness, apes have an appre­ci­a­tion of the oth­er’s sit­u­a­tion and show a degree of per­spec­tive-tak­ing. One strik­ing report in this regard con­cerns a bonobo female named Kuni, who found a wound­ed bird in her enclo­sure at Twycross Zoo, in Eng­land. Kuni picked up the bird, and when her keep­er urged her to let it go, she climbed to the high­est point of the high­est tree, care­ful­ly unfold­ed the bird’s wings and spread them wide open, one wing in each hand, before throw­ing it as hard as she could toward the bar­ri­er of the enclo­sure. When the bird fell short, Kuni climbed down and guard­ed it until the end of the day, when it flew to safe­ty. Obvi­ous­ly, what Kuni did would have been inap­pro­pri­ate toward a mem­ber of her own species. Hav­ing seen birds in flight many times, she seemed to have a notion of what would be good for a bird, thus giv­ing us an anthro­poid illus­tra­tion of Smith’s “chang­ing places in fancy.”

This is not to say that all we have are anec­dotes. Sys­tem­at­ic stud­ies have been con­duct­ed on so-called “con­so­la­tion” behav­ior. Con­so­la­tion is defined as friend­ly or reas­sur­ing behav­ior by a bystander toward a vic­tim of aggres­sion. For exam­ple, chim­panzee A attacks chim­panzee B, after which bystander C comes over and embraces or grooms B. Based on hun­dreds of such obser­va­tions, we know that con­so­la­tion occurs reg­u­lar­ly and exceeds base­line lev­els of con­tact. In oth­er words, it is a demon­stra­ble ten­den­cy that prob­a­bly reflects empa­thy, since the objec­tive of the con­sol­er seems to be to alle­vi­ate the dis­tress of the oth­er. In fact, the usu­al effect of this kind of behav­ior is that it stops scream­ing, yelp­ing, and oth­er signs of distress.

A bot­tom-up view of empathy 

The above exam­ples help explain why to the biol­o­gist, a Russ­ian doll is such a sat­is­fy­ing play­thing, espe­cial­ly if it has a his­tor­i­cal dimen­sion. I own a doll of Russ­ian Pres­i­dent Vladimir Putin, with­in whom we dis­cov­er Yeltsin, Gor­bachev, Brezh­nev, Kruschev, Stal­in, and Lenin, in that order. Find­ing a lit­tle Lenin and Stal­in with­in Putin will hard­ly sur­prise most polit­i­cal ana­lysts. The same is true for bio­log­i­cal traits: The old always remains present in the new.

This is rel­e­vant to the debate about the ori­gins of empa­thy, espe­cial­ly because of the ten­den­cy in some dis­ci­plines, such as psy­chol­o­gy, to put human capac­i­ties on a pedestal. They essen­tial­ly adopt a top-down approach that empha­sizes the unique­ness of human lan­guage, con­scious­ness, and cog­ni­tion. But instead of try­ing to place empa­thy in the upper regions of human cog­ni­tion, it is prob­a­bly best to start out exam­in­ing the sim­plest pos­si­ble process­es, some per­haps even at the cel­lu­lar lev­el. In fact, recent neu­ro­science research sug­gests that very basic process­es do under­lie empa­thy. Researchers at the Uni­ver­si­ty of Par­ma, in Italy, were the first to report that mon­keys have spe­cial brain cells that become active not only if the mon­key grasps an object with its hand but also if it mere­ly watch­es anoth­er do the same. Since these cells are acti­vat­ed as much by doing as by see­ing some­one else do, they are known as mir­ror neu­rons, or “mon­key see, mon­key do” neurons.

It seems that devel­op­men­tal­ly and evo­lu­tion­ar­i­ly, advanced forms of empa­thy are pre­ced­ed by and grow out of more ele­men­tary ones. Biol­o­gists pre­fer such bot­tom-up accounts. They always assume con­ti­nu­ity between past and present, child and adult, human and ani­mal, even between humans and the most prim­i­tive mammals.

So, how and why would this trait have evolved in humans and oth­er species? Empa­thy prob­a­bly evolved in the con­text of the parental care that char­ac­ter­izes all mam­mals. Sig­nal­ing their state through smil­ing and cry­ing, human infants urge their care­giv­er to take action. This also applies to oth­er pri­mates. The sur­vival val­ue of these inter­ac­tions is evi­dent from the case of a deaf female chim­panzee I have known named Krom, who gave birth to a suc­ces­sion of infants and had intense pos­i­tive inter­est in them. But because she was deaf, she would­n’t even notice her babies’ calls of dis­tress if she sat down on them. Krom’s case illus­trates that with­out the prop­er mech­a­nism for under­stand­ing and respond­ing to a child’s needs, a species will not survive.

Dur­ing the 180 mil­lion years of mam­malian evo­lu­tion, females who respond­ed to their off­spring’s needs out-repro­duced those who were cold and dis­tant. Hav­ing descend­ed from a long line of moth­ers who nursed, fed, cleaned, car­ried, com­fort­ed, and defend­ed their young, we should not be sur­prised by gen­der dif­fer­ences in human empa­thy, such as those pro­posed to explain the dis­pro­por­tion­ate rate of boys affect­ed by autism, which is marked by a lack of social com­mu­ni­ca­tion skills.

Empa­thy also plays a role in coop­er­a­tion. One needs to pay close atten­tion to the activ­i­ties and goals of oth­ers to coop­er­ate effec­tive­ly. A lioness needs to notice quick­ly when oth­er lioness­es go into hunt­ing mode, so that she can join them and con­tribute to the pride’s suc­cess. A male chim­panzee needs to pay atten­tion to his bud­dy’s rival­ries and skir­mish­es with oth­ers so that he can help out when­ev­er need­ed, thus ensur­ing the polit­i­cal suc­cess of their part­ner­ship. Effec­tive coop­er­a­tion requires being exquis­ite­ly in tune with the emo­tion­al states and goals of others.

With­in a bot­tom-up frame­work, the focus is not so much on the high­est lev­els of empa­thy, but rather on its sim­plest forms, and how these com­bine with increased cog­ni­tion to pro­duce more com­plex forms of empa­thy. How did this trans­for­ma­tion take place? The evo­lu­tion of empa­thy runs from shared emo­tions and inten­tions between indi­vid­u­als to a greater self/other distinction—that is, an “unblur­ring” of the lines between indi­vid­u­als. As a result, one’s own expe­ri­ence is dis­tin­guished from that of anoth­er per­son, even though at the same time we are vic­ar­i­ous­ly affect­ed by the oth­er’s. This process cul­mi­nates in a cog­ni­tive appraisal of the oth­er’s behav­ior and sit­u­a­tion: We adopt the oth­er’s perspective.

As in a Russ­ian doll, how­ev­er, the out­er lay­ers always con­tain an inner core. Instead of evo­lu­tion hav­ing replaced sim­pler forms of empa­thy with more advanced ones, the lat­ter are mere­ly elab­o­ra­tions on the for­mer and remain depen­dent on them. This also means that empa­thy comes nat­u­ral­ly to us. It is not some­thing we only learn lat­er in life, or that is cul­tur­al­ly con­struct­ed. At heart, it is a hard-wired response that we fine-tune and elab­o­rate upon in the course of our lives, until it reach­es a lev­el at which it becomes such a com­plex response that it is hard to rec­og­nize its ori­gin in sim­pler respons­es, such as body mim­ic­ry and emo­tion­al contagion.

On a leash

Biol­o­gy holds us “on a leash,” in the felic­i­tous words of biol­o­gist Edward Wil­son, and will let us stray only so far from who we are. We can design our life any way we want, but whether we will thrive depends on how well that life fits human predispositions.

I hes­i­tate to pre­dict what we humans can and can’t do, but we must con­sid­er our bio­log­i­cal leash when decid­ing what kind of soci­ety we want to build, espe­cial­ly when it comes to goals like achiev­ing uni­ver­sal human rights.

If we could man­age to see peo­ple on oth­er con­ti­nents as part of us, draw­ing them into our cir­cle of reci­procity and empa­thy, we would be build­ing upon, rather than going against, our nature.

For instance, in 2004, the Israeli Min­is­ter of Jus­tice caused polit­i­cal uproar for sym­pa­thiz­ing with the ene­my. Yosef Lapid ques­tioned the Israeli army’s plans to demol­ish thou­sands of Pales­tin­ian homes in a zone along the Egypt­ian bor­der. He had been touched by images on the evening news. “When I saw a pic­ture on the TV of an old woman on all fours in the ruins of her home look­ing under some floor tiles for her med­i­cines, I did think, ‘What would I say if it were my grand­moth­er?’ ” he said. Lapid’s grand­moth­er was a Holo­caust victim.

This inci­dent shows how a sim­ple emo­tion can widen the def­i­n­i­tion of one’s group. Lapid had sud­den­ly real­ized that Pales­tini­ans were part of his cir­cle of con­cern, too. Empa­thy is the one weapon in the human reper­toire that can rid us of the curse of xenophobia.

Empa­thy is frag­ile, though. Among our close ani­mal rel­a­tives, it is switched on by events with­in their com­mu­ni­ty, such as a young­ster in dis­tress, but it is just as eas­i­ly switched off with regards to out­siders or mem­bers of oth­er species, such as prey. The way a chim­panzee bash­es in the skull of a live mon­key by hit­ting it against a tree trunk is no adver­tise­ment for ape empa­thy. Bono­bos are less bru­tal, but in their case, too, empa­thy needs to pass through sev­er­al fil­ters before it will be expressed. Often, the fil­ters pre­vent expres­sions of empa­thy because no ape can afford feel­ing pity for all liv­ing things all the time. This applies equal­ly to humans. Our evo­lu­tion­ary back­ground makes it hard to iden­ti­fy with out­siders. We’ve evolved to hate our ene­mies, to ignore peo­ple we bare­ly know, and to dis­trust any­body who does­n’t look like us. Even if we are large­ly coop­er­a­tive with­in our com­mu­ni­ties, we become almost a dif­fer­ent ani­mal in our treat­ment of strangers.

This is the chal­lenge of our time: glob­al­iza­tion by a trib­al species. In try­ing to struc­ture the world such that it suits human nature, the point to keep in mind is that polit­i­cal ide­o­logues by def­i­n­i­tion hold nar­row views. They are blind to what they don’t wish to see. The pos­si­bil­i­ty that empa­thy is part of our pri­mate her­itage ought to make us hap­py, but we are not in the habit of embrac­ing our nature. When peo­ple kill each oth­er, we call them “ani­mals.” But when they give to the poor, we praise them for being “humane.” We like to claim the lat­ter ten­den­cy for our­selves. Yet, it will be hard to come up with any­thing we like about our­selves that is not part of our evo­lu­tion­ary back­ground. What we need, there­fore, is a vision of human nature that encom­pass­es all of our ten­den­cies: the good, the bad, and the ugly.

Our best hope for tran­scend­ing trib­al dif­fer­ences is based on the moral emo­tions, because emo­tions defy ide­ol­o­gy. In prin­ci­ple, empa­thy can over­ride every rule about how to treat oth­ers. When Oskar Schindler kept Jews out of con­cen­tra­tion camps dur­ing World War II, for exam­ple, he was under clear orders by his soci­ety on how to treat peo­ple, yet his feel­ings interfered.

Car­ing emo­tions may lead to sub­ver­sive acts, such as the case of a prison guard who dur­ing wartime was direct­ed to feed his charges only water and bread, but who occa­sion­al­ly sneaked in a hard-boiled egg. How­ev­er small his ges­ture, it etched itself into the pris­on­ers’ mem­o­ries as a sign that not all of their ene­mies were mon­sters. And then there are the many acts of omis­sion, such as when sol­diers could have killed cap­tives with­out neg­a­tive reper­cus­sions but decid­ed not to. In war, restraint can be a form of compassion.

Emo­tions trump rules. This is why, when speak­ing of moral role mod­els, we talk of their hearts, not their brains (even if, as any neu­ro­sci­en­tist will point out, the heart as the seat of emo­tions is an out­dat­ed notion). We rely more on what we feel than what we think when solv­ing moral dilemmas.

It’s not that reli­gion and cul­ture don’t have a role to play, but the build­ing blocks of moral­i­ty clear­ly pre­date human­i­ty. We rec­og­nize them in our pri­mate rel­a­tives, with empa­thy being most con­spic­u­ous in the bonobo ape and reci­procity in the chim­panzee. Moral rules tell us when and how to apply our empath­ic ten­den­cies, but the ten­den­cies them­selves have been in exis­tence since time immemorial.

Frans B. M. de Waal, Ph.D., a Dutch-born pri­ma­tol­o­gist, is the C. H. Can­dler Pro­fes­sor at Emory Uni­ver­si­ty and direc­tor of the Liv­ing Links Cen­ter at the Yerkes Nation­al Pri­mate Research Cen­ter in Atlanta. This essay is adapt­ed from his book, Our Inner Ape: A Lead­ing Pri­ma­tol­o­gist Explains Why We Are Who We Are. His lat­est book is The Age of Empa­thy. Greater Good Mag­a­zine, based at UC-Berke­ley, is a quar­ter­ly mag­a­zine that high­lights ground break­ing sci­en­tif­ic research into the roots of com­pas­sion and altruism.

Relat­ed arti­cles by Greater Good Magazine:


  1. John on February 3, 2010 at 7:40

    Very inter­est­ing arti­cle, did­n’t know much about Empa­thy before read­ing this and I def­i­nite­ly learned a lot.

About SharpBrains

SHARPBRAINS is an independent think-tank and consulting firm providing services at the frontier of applied neuroscience, health, leadership and innovation.
SHARPBRAINS es un think-tank y consultoría independiente proporcionando servicios para la neurociencia aplicada, salud, liderazgo e innovación.

Top Articles on Brain Health and Neuroplasticity

Top 10 Brain Teasers and Illusions


Subscribe to our e-newsletter

* indicates required

Got the book?